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Climate, satellite imagery and the seasonal abundance of the tick Rhipicephalus appendiculatus in southern Africa: a new perspective.

Recent predictive models for the distribution of the African tick Rhipicephalus appendiculatus Neumann, based on the computer packages CLIMEX and BIOCLIM and data derived from meteorological satellites, and for the seasonal dynamics of the same tick using the computer simulation models ECFXPERT and T3HOST, all have their limitations. Statistical analysis of the relationships between the seasonal abundance of all three life stages of this tick and climatic and satellite-derived data from five sites in southern Africa, taken from the literature, supports a new perspective that it is the timing of the questing activity of the desiccation-vulnerable larvae that determines the pattern of the tick's seasonal dynamics. The timing of the activity of nymphs and adults is determined by temperature-dependent development rates plus the delaying phenomenon of photoperiod-sensitive diapause, the timing and duration of which have evolved to achieve maximum generation survival by ensuring the occurrence of eggs and larvae during periods of optimal climatic conditions. The most important environmental factor appears to be night-time minimum temperature, determining condensation and saturation deficit and thus the tick's ability to replenish moisture lost during the daytime and so to survive while questing for hosts. It is the larvae whose numbers are correlated most closely with these factors, consistent with earlier experimental results showing larvae to be most susceptible to desiccating conditions. There is a statistical linkage between larval tick numbers and satellite imagery, arising from the correlation between larval numbers and minimum temperature and saturation deficit conditions, and in turn the relationship between these climatic conditions and the subsequent vegetational changes monitored by the satellites. Moisture availability to larvae is likely to be the critical factor throughout the geographical range of R. appendiculatus, but the precise combination of climatic conditions that optimize moisture availability and questing tick survival can be expected to vary geographically. The relationships between ticks, temperatures and satellite data in parts of equatorial Africa have yet to be established. These correlative patterns highlight both the critical life stage and environmental factors when trying to understand temporal, and ultimately spatial, variations in tick abundance.

Animals↗

Effect of prior exposure to noninfected ticks on susceptibility of mice to Lyme disease spirochetes.

To determine whether prior exposure to Nearctic Ixodes vector ticks protects native reservoir mice from tick-borne infection by Lyme disease spirochetes, we compared their infectivities for white-footed mice and laboratory mice that had been repeatedly infested by noninfected deer ticks. Nymphal ticks readily engorged on tick-exposed laboratory mice, but their feeding success on white-footed mice progressively declined. Tick-borne spirochetes readily infected previously tick-infested mice. Thus, prior infestation by Nearctic ticks does not protect sympatric reservoir mice or Palearctic laboratory mice from infection by sympatric tick-borne spirochetes.

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Ehrlichia phagocytophila genogroup rickettsiae in ixodid ticks from California collected in 1995 and 1996.

A total of 1,246 ixodid ticks collected in 1995 and 1996 from seven California counties were examined for the presence of Ehrlichia phagocytophila genogroup rickettsiae by using a nested PCR technique. Of 1,112 adult Ixodes pacificus Cooley and Kohls ticks tested, nine pools, each containing five ticks, were positive (minimum percentage of ticks harboring detectable ehrlichiae, 0.8%). Positive ticks were limited to four of the seven counties (Sonoma, El Dorado, Santa Cruz, and Orange). In Santa Cruz County, three positive pools were identified at the home of an individual with prior confirmed human granulocytic ehrlichiosis. In El Dorado County, positive ticks were found at sites where cases of granulocytic ehrlichiosis in a horse and a llama had recently occurred. Among 47 nymphal I. pacificus ticks collected in Sonoma County, one positive pool was identified. Fifty-seven adult Dermacentor occidentalis Marx and 30 adult D. variabilis Say ticks, collected chiefly in southern California, were negative. These data, although preliminary, suggest that the prevalence of E. phagocytophila genogroup rickettsiae in ixodid ticks of California may be lower than in cognate vector populations (i.e., I. scapularis Say = I. dammini Spielman, Clifford, Piesman, and Corwin) in the eastern and midwestern United States.

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Invasion and intracellular development of the human granulocytic ehrlichiosis agent in tick cell culture.

Human granulocytotropic ehrlichias are tick-borne bacterial pathogens that cause an acute, life-threatening illness, human granulocytic ehrlichiosis (HGE). Ehrlichias within neutrophil granulocytes that invade tick bite sites are likely ingested by the vector, to be transmitted to another mammalian host during the tick's next blood meal. Thus, the cycle of replication and development in the vector is prerequisite to mammalian infection, and yet these events have not been described. We report tick cell culture isolation of two strains of the HGE agent directly from an infected horse and a dog and have also established a human isolate from HL60 culture in tick cells, proving that the blood stages of the HGE agent are infectious for tick cells, as are those replicating in the human cell line HL60. This required changes to the culture system, including a new tick cell line. In tick cell layers, the HGE agent induced foci of infection that caused necrotic plaques and eventual destruction of the culture. Using the human isolate and electron microscopy, we monitored adhesion, internalization, and replication in vector tick cells. Both electron-lucent and -dense forms adhered to and entered cells by a mechanism reminiscent of phagocytosis. Ehrlichial cell division was initiated soon after, resulting in endosomes filled with numerous ehrlichias. During early development, pale ehrlichias with a tight cell wall dominated, but by day 2, individual bacteria condensed into dark forms with a rippled membrane. These may become compacted into clumps where individual organisms are barely discernible. Whether these are part of an ehrlichia life cycle or are degenerating is unknown.

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Roles of neutrophil beta 2 integrins in kinetics of bacteremia, extravasation, and tick acquisition of Anaplasma phagocytophila in mice.

Tick saliva contains anti-inflammatory and immunosuppressive substances that facilitate blood feeding and enhance tick-vectored pathogen transmission, including Anaplasma phagocytophila an etiologic agent of granulocytic ehrlichiosis. As such, inflammation at a tick-feeding site is strikingly different than that typically observed at other sites of inflammation. Up-regulation of CD11b/CD18 occurs in host granulocytes following interaction or infection with A phagocytophila, and the absence of CD11b/CD18 results in early increases in bacteremia. We hypothesized that beta 2 integrin-dependent infection kinetics and leukocyte extravasation are important determinants of neutrophil trafficking to, and pathogen acquisition at, tick-feeding sites. A phagocytophila infection kinetics were evaluated in CD11a/CD18, CD11b/CD18, and CD18 knock-out mice using quantitative polymerase chain reaction (PCR) of blood, ticks, and skin biopsies in conjunction with histopathology. A marked increase in the rate of A phagocytophila infection of neutrophils and pathogen burden in blood followed tick feeding. Infection kinetics were modified by beta 2 integrin expression and systemic neutrophil counts. Significant neutrophil-pathogen trafficking was observed to both suture and tick sites. Despite the prominent role for beta 2 integrins in neutrophil arrest in flowing blood, successful pathogen acquisition by ticks occurred in the absence of beta 2 integrins. Establishment of feeding pools that rely less on leukocyte trafficking and more on small hemorrhages may explain the ready amplification of A phagocytophila DNA from ticks infested on CD11/CD18-deficient mouse strains.

Anaplasma phagocytophilum↗

Reduced incidence of Babesia bigemina infection in cattle immunized against the cattle tick, Boophilus microplus.

Boophilus microplus is an important vector of bovine disease agents having a major economic impact on cattle production in many tropical and subtropical countries. Components of tick saliva that enable ticks to feed may also facilitate establishment of tick-borne pathogens in the vertebrate host. It has been suggested that acquired resistance against molecules in tick saliva could inhibit parasite transmission, and there is increasing evidence to support this hypothesis. The effect of immune resistance to B. microplus on the incidence of tick-transmitted pathogens was the focus of this experiment. Groups of four dairy cows were injected with antigen extracts of tick salivary glands, midgut, adjuvant only, or PBS, prior to a grazing period in a pasture in Thailand where ticks are abundant and babesiosis is enzootic. These animals were then observed for evidence of babesiosis throughout the rainy season. A reduction in the incidence of clinical babesiosis was observed among cattle immunized with salivary gland preparations compared to nonimmunized controls (P < 0.05). Immunization with midgut or adjuvant only both resulted in a slight reduction in observed disease compared to the same negative control group. B. bigemina was detected in fewer ticks (24.43%) collected from salivary gland-immunized cattle than those collected from the remaining groups (> or =44.57%). These results indicated that immunization with salivary gland antigens could affect pathogen transmission and appears promising for control of tick-borne diseases of cattle.

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Natural infection, transovarial transmission, and transstadial survival of Rickettsia bellii in the Tick Ixodes loricatus (Acari: Ixodidae) from Brazil.

An Ixodes loricatus engorged female, infected with Rickettsia bellii, was collected from an opossum (Didelphis aurita) in Mogi das Cruzes, São Paulo State, Brazil. Two consecutive laboratory tick generations (F(1) and F(2)) reared from this single engorged female were evaluated for Rickettsia infection by polymerase chain reaction (PCR) targeting specific Rickettsia genes. Immature ticks fed on naïve Wistar rats (Rattus norvegicus) and adult ticks fed on opossum (D. aurita), both free of ticks and rickettsial infection. PCR performed on individual ticks from the F(1) (20 larvae, 10 nymphs, and 10 adults) and the F(2) (30 larvae, 30 nymphs, and 15 adults) yielded expected bands compatible with Rickettsia. All the PCR products that were sequenced, targeting gltA gene, resulted in sequences identical to each other and 99.7% (349/350) similar to the corresponding sequence of R. bellii in GenBank. The R. bellii infection on ticks from the second laboratory generation (F(2)) was confirmed by other PCR protocols and successful isolation of R. bellii in cell culture. We report for the first time a Rickettsia species infecting I. loricatus, and the first report of R. bellii in the tick genus Ixodes. We conclude that there was an efficient transovarial transmission and transstadial survival of this Rickettsia species in the tick I. loricatus. Our results suggest that R. bellii might be maintained in nature solely by transovarial transmission and transstadial survival in ticks (no amplifier vertebrate host is needed), since there has been no direct or indirect evidence of infection of vertebrate hosts by R. bellii.

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Prevalence of spotted fever group Rickettsia species detected in ticks in La Rioja, Spain.

Our objective was to learn the prevalence of spotted fever group (SFG) Rickettsia detected in ticks in La Rioja, in the north of Spain. From 2001 to 2005, 496 ticks representing 7 tick species were analysed at the Hospital de La Rioja. Ticks were removed from humans with or without rickettsial syndrome (n = 59) or collected from mammals (n = 371) or from vegetation by dragging (n = 66). The presence of SFG Rickettsia in these ticks was investigated by semi-nested PCR (ompA gene) and sequencing. A phylogenetic tree using Clustal method (neighbor-joining) was constructed with these data. Only 3 of 170 Hyalomma marginatum ticks carried SFG Rickettsia. Sequencing analysis demonstrated the presence of Rickettsia aeschlimannii (1.8%). Furthermore, Rickettsia massiliae and BAR29 were found in 3 of 120 Rhipicephalus sanguineus specimens (2.5%). In contrast, 81 of 83 tested Dermacentor marginatus ticks were PCR-positive (97%). Rickettsia slovaca (40.6%) and Rickettsia sp. strains RpA4, DnS14, DnS28 and JL-02 (59.3%) were found within this tick species. No SFG Rickettsia was detected using ompA primers when Ixodes ricinus, Rhipicephalus bursa, Rhipicephalus turanicus, Rhipicephalus eversti eversti, Hyalomma detritum scupense and Rhipicephalus sp. were analyzed. We detected 17.5% of ticks associated with different SFG Rickettsia: R. aeschlimannii, R. massiliae, BAR29, R. slovaca and Rickettsia sp. strains RpA4, DnS14, DnS28 and JL-02. Their presence has to be taken into account since most of them have been recognized as human pathogens.

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Reduced abundance of Ixodes scapularis (Acari: Ixodidae) and the tick parasitoid Ixodiphagus hookeri (Hymenoptera: Encyrtidae) with reduction of white-tailed deer.

The principal vector for the pathogens of Lyme disease, human granulocytic ehrlichiosis, and human babesiosis is the tick Ixodes scapularis Say. A chalcid wasp, Ixodiphagus hookeri, in the family Encyrtidae parasitizes populations of the tick on several islands or other geographically isolated sites in New England with high densities of these ticks and white-tailed deer (Odocoileus virginianus), the principal host for adult I. scapularis. Deer densities were reduced at a forested tract in Bridgeport and the Bluff Point Coastal Reserve in Groton, Connecticut, from levels exceeding 90 animals per km2 in 1992 (Bridgeport) and 1994 (Bluff Point) to 17 and 10 animals per km2, respectively, by fall 2001. Tick densities declined with sustained reductions in the population of white-tailed deer. Similarly, prevalence of tick parasitism by Ixodes hookeri declined at both sites from 30 to 25% to <1.0% and was significantly correlated with previous year's deer density at both sites (r(s) = 0.933 and r(s) = 0.867, P < or = 0.0001) and with nymphal tick densities at Bridgeport (r(s) = 0.867, P < or = 0.0001), but was not as well correlated with tick densities in Groton. The virtual disappearance of I. hookeri in this study corresponds with a lack of I. hookeri in mainland I. scapularis at comparable deer and tick densities, suggesting that there is a threshold deer density of approximatley 10-20/km2, with corresponding tick densities necessary for I. hookeri to successfully parasitize I. scapularis.

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Incidence of tick-bite in man in Aland Islands: reference to the spread of Lyme borreliosis.

The common tick Ixodes ricinus is a vector both for the virus of tick-borne viral encephalitis (Kumlinge disease, KD) and of Borrelia burgdorferi (Lyme borreliosis, LB). Bites of the tick are believed to be common in Aland, an island province of Finland in the Baltic Sea. KD has been seen there for many years, and cases of LB have been diagnosed lately. The purpose of this study was to find the incidence of tick-bite among the population with possible implications for the spread of LB. A questionnaire was sent to 561 persons greater than 8 years of age out of a total of 21,281 in order to record tick-bites and erythema chronicum migrans (ECM). The number of answers was 519 (92.5%). 441 persons (85%) had been bitten by ticks, 146 greater than 10 times. 14 probands had had ECM, 73 other rashes around the tick-bite. It is concluded that tick-bites are very common among the population, and thus conditions are favourable for the spread of any tick-borne infection such as LB. The number of patients with secondary or tertiary LB diagnosed in the area is presently 17. It is likely that there are many undetected cases in the area.

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Preliminary survey of ticks (Acari: Ixodidae) on cattle in northern Sudan.

In a cross sectional survey conducted during the period June 2001 to July 2002, the geographical distribution of ticks on cattle in the Sudan was determined. Seventeen locations were surveyed from Northern, Central, Eastern, Western, Blue Nile and White Nile Provinces. Total body collections of ticks were made from 20 cattle at each location. Four tick genera and 11 species were identified. The tick species collected included Amblyomma lepidum, Amblyomma variegatum, Boophilus decoloratus, Hyalomma anatolicum anatolicum, Hyalomma dromedarii, Hyalomma impeltatum, Hyalomma marginatum rufipes, Hyalomma truncatum, Rhipicephalus evertsi evertsi, Rhipicephalus sanguineus group and Rhipicephalus simus simus. Major ecological changes have occurred due to extensive animal movement, deforestation, desertification and establishment of large mechanized agricultural schemes. These factors have certainly affected the distribution of ticks and tick-borne diseases in the Sudan. The absence of A. variegatum and A. lepidum in northern Sudan was not surprising, since these tick species are known to survive in humid areas and not in the desert and semi-desert areas of northern Sudan. The absence of B. annulatus in northern and central Sudan is in accordance with the finding that this tick species is restricted to the southern parts of the central Sudan. The presence of H. anatolicum anatolicum in Um Benin in relatively high abundance is an interesting finding. The present finding may indicate that the southern limit of this species has changed and moved southwards to latitude 13 degrees N. It is concluded that major changes in tick distribution have taken place in the Sudan.

Animal Husbandry↗

Clustering of host-seeking nymphal deer ticks (Ixodes dammini) infected by Lyme disease spirochetes (Borrelia burgdorferi).

In areas where the agent of Lyme disease is intensely enzootic, the mouse reservoirs may be universally infected. Because a large proportion of the vector tick population appears to feed upon these hosts, the prevalence of infection in the vectors should approach 100%. However, infection in host-seeking nymphal ticks in nature rarely exceeds 40%. To help reconcile this apparent paradox, we examined whether estimates of prevalence might differ if we did not assume that infected ticks are randomly or uniformly distributed within a site. Nymphal Ixodes dammini were collected by dragging a series of 10-meter replicates within an intensely enzootic site. Estimates of the prevalence of spirochetal infection, based upon the monthly means of individual 10-meter collections, were then compared with estimates derived by pooling all samples. Host-seeking ticks tended to cluster. The Lyme disease spirochete was present in 15.6% of 469 pooled ticks. When the prevalence estimate was based solely on ticks in clusters that contained one or more infected ticks, however, at least 50% of the ticks were infected. We conclude that nymphal deer ticks infected by Lyme disease spirochetes tend to aggregate spatially in nature, and that prevalence estimates based upon a mean value for pools may be misleading.

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Factors influencing the distribution of larval blacklegged ticks on rodent hosts.

Because of differences among hosts in reservoir competence for tick-borne diseases, the distribution of larval blacklegged ticks on hosts might determine tick infection prevalence and disease risk to humans. We conducted a three-part study to determine the factors responsible for greater burdens of larval blacklegged ticks on white-footed mice than on eastern chipmunks. A microhabitat study indicated that questing ticks have higher encounter rates with mice than with chipmunks. Laboratory experiments demonstrated that ticks oriented more strongly toward mice. However, larval ticks fed more successfully from chipmunks. Our results strongly suggest that mice are both more likely to use larval tick-infested microhabitats and to attract questing larvae than are chipmunks, leading to a dramatically higher initial infestation rate, which is then reduced by greater grooming activity by mice. The high mortality rate of larvae that were experimentally introduced onto mice suggests that grooming is a significant cause of mortality to larval blacklegged ticks.

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[Risk of arbovirus transmission by ticks in Thailand].

Ticks are known vectors of transmission for a number of infectious viral diseases from wild or domestic animals to humans. Many tick-borne diseases cause severe clinical syndromes such as encephalitis or hemorrhagic fever. Animal carriers of enzootic diseases are often asymptomatic. Within the framework of a program to monitor emerging viral diseases in Southeast Asia, identification of ticks capable of transmitting diseases from animals to man is a prerequisite for epidemiologic study to assess the risk of tick-borne disease. The purpose of this report is to provide an update on the situation based on a study of tick fauna in the central plain area of Thailand and on current knowledge about tick-borne arboviruses in Asia. In addition to an exhaustive inventory of ticks found in Thailand, this study describes the viruses with pathogenic potential that ticks are known to carry and transmit to man. A perusal of the literature allowed initial assessment of the risk for introduction and spread of ticks by bird hosts in Southeast Asia as well as of associated virus.

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Studies on the occurrence of Gram-negative bacteria in ticks: Ixodes ricinus as a potential vector of Pasteurella.

A total of 372 Ixodes ricinus ticks (101 females, 122 males, and 149 nymphs) collected by flagging in 6 mixed woodlands of eastern Poland were examined by culture for the presence of internal Gram-negative bacteria other than Borrelia burgdorferi. Adult ticks were examined in pools of 2 specimens each and nymphs were examined in pools of 3-5 specimens each. Ticks were disinfected in 70 % ethanol and homogenized in 0.85% NaCl. The diluted homogenate was inoculated onto 3 kinds of agar media: buffered charcoal yeast extract (BCYE-alpha) for isolation of fastidious Gram-negative bacteria, eosin methylene blue agar (EMB) for isolation of enterobacteria, and tryptic soya agar for isolation of all other non-fastidious Gram-negative bacteria. The Gram-negative isolates were identified with the API Systems 20E and NE microtests. A total of 9 species of Gram-negative bacteria were identified, of which the commonest were strains determined as Pasteurella pneumotropica/haemolytica, which were isolated on BCYE-alpha agar from ticks collected in all 6 examined woodlands. The total number of these strains (49) exceeded the total number of all other strains of Gram-negative bacteria recovered from ticks (30). Of the total number of examined ticks, the minimum infection rate with Pasteurella pneumotropica/haemolytica was highest in females (18.8%), and slightly lower in males (12.3%) and nymphs (10%). Besides Pasteurella pneumotropica/haemolytica, the following species of Gram-negative bacteria were isolated from examined ticks: Pantoea agglomerans, Serratia marcescens, Serratia plymuthica on EMB agar and Aeromonas hydrophila, Burkholderia cepacia, Chromobacterium violaceum, Pseudomonas aeruginosa, and Stenotrophomonas maltophilia on tryptic soya agar. Minimal infection rates with these bacteria were low, ranging from 0.7-5.9%. Of the isolated bacteria, Chromobacterium violaceum, Pasteurella pneumotropica/haemolytica, Pseudomonas aeruginosa, and Serratia marcescens are potentially pathogenic for man and/or animals. In particular, the common occurrence of Pasteurella pneumotropica/haemolytica in Ixodes ricinus ticks poses a potential risk of pasteurellosis for humans and animals exposed to tick bites.

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Occurrence of hard ticks in dogs from Warsaw area.

Ticks are known as a vector of many viral, bacterial, rickettsial and protozoan infections of dogs. The threat with particular tick-transmitted disease depends very much on the tick species which feeds on the dog. In this paper we present results of research designed to identify tick species which attack dogs in the Warsaw area. Ticks were collected from dogs presented for medical examination in veterinary clinics in different parts of Warsaw. All of 590 tick specimens found on dogs belonged to the family Ixodidae (hard ticks). The majority (64.6 %) of the collected ticks were identified as Dermacentor reticulatus--vector of Babesia canis. The other tick species found on examined dogs was Ixodes ricinus. The male to female ratio was more than 3 times higher in D. reticulatus than in I. ricinus.

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Cutaneous basophil responses and immune resistance of guinea pigs to ticks: passive transfer with peritoneal exudate cells or serum.

Resistance to infestation by larval Amblyomma americanum or Rhipicephalus sanguineus ticks was transferred to naive guinea pigs with peritoneal exudate cells (PEC) or serum from donors immunized by prior infestation with homologous tick larvae. In the A. americanum system, PEC transfer induced 87% tick rejection, which was similar to the level of resistance in actively sensitized hosts. In the R. sanguineus system, PEC conferred resistance (39% rejection) that was weaker than in actively sensitized hosts (57% rejection). In both systems, immune serum conferred significant but weaker resistance (20 to 29% rejection). In actively sensitized hosts, resistance to each tick species was specific, but there was considerable cross-reactive resistance. Basophils dominated the 24-hr challenge feeding sites of A. americanum ticks in actively sensitized hosts (69% of the infiltrate) and recipients of sensitized PEC (69%). Mononuclear cells were dominant (69% of the infiltrate) in the challenged tissues of immune serum recipients that had a significant but weaker cutaneous basophil response (24%). Mononuclear cells also dominated (58% of the infiltrate) the 24-hr challenge feeding sites of R. sanguineus ticks in actively sensitized hosts, but there were also 24% basophils. These studies demonstrate that immune resistance to tick is dependent on sensitized lymphoid cells or serum components, and that sensitized cells or serum can transfer a cutaneous basophil response that is associated with immune resistance. Rejection of ticks is usually associated with large basophil infiltrates, but sometimes mononuclear cells are dominant. Thus, immune resistance of guinea pigs to ticks is a heterogeneous response in which immune cells and serum probably act to recruit diverse effector leukocytes to mediate rejection that is specific but significantly cross-reactive.

Animals↗

Mechanisms of eosinophilia in mice infested with larval Haemaphysalis longicornis ticks.

Infestation of larval Haemaphysalis longicornis ticks induced a threefold increase of eosinophils in the peripheral blood of normal WBB6F1- +/+ mice 2 days after tick infestation. In genetically mast cell-deficient WBB6F1- W/Wv mice, a threefold increase of blood eosinophils was observed 6 days after the tick infestation. However, marked infiltration of eosinophils was detected in the tick infestation sites of the WBB6F1- +/+ mice but not the WBB6F1- W/Wv mice. When the mast cell deficiency of WBB6F1- W/Wv mice had been rescued locally by intradermal injections of WBB6F1- +/+ mouse-derived cultured mast cells, a rapid increase of blood eosinophils and tissue infiltration of eosinophils were revealed following tick infestation. The intravenous (i.v.) injection of immune spleen or lymph node cells obtained from WBB6F1- +/+ mice 10 days after tick infestation led to significant eosinophilia in naive recipient mice. Treatment with anti-Thy-1.2 or anti-CD4 monoclonal antibody (mAb) and complement (C) completely abolished the eosinophilia; the early response (2 days after tick challenge) is dependent on mast cells at the feeding site, and the late response (6 days after tick challenge) is dependent on T lymphocytes. Since amplified interleukin-5 (IL-5) cDNA was detectable in the spleen cells 4 days after tick infestation, the late response might be mediated by IL-5. The infiltration of eosinophils at the feeding site of skin appeared to be dependent on mast cells.

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